Unintended consequences of selection for increased production on the health and welfare of livestock
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Review
Arch. Anim. Breed., 64, 177–185, 2021
Open Access
https://doi.org/10.5194/aab-64-177-2021
© Author(s) 2021. This work is distributed under
Archives Animal Breeding
the Creative Commons Attribution 4.0 License.
Unintended consequences of selection for increased
production on the health and welfare of livestock
Este van Marle-Köster and Carina Visser
Faculty of Natural and Agricultural Sciences, Department of Animal Science,
University of Pretoria, Pretoria 0028, South Africa
Correspondence: Este van Marle-Köster (evm.koster@up.ac.za)
Received: 2 November 2020 – Revised: 24 March 2021 – Accepted: 13 April 2021 – Published: 25 May 2021
Abstract. Modern farming technologies, including quantitative selection and breeding methods in farm animal
species, resulted in increased production and efficiency. Selection for increased output in both intensive and ex-
tensive production systems has trade-offs and negative outcomes, often more pronounced in intensive systems.
Animal welfare and health are often adversely affected and this influences sustainable production. The relative
importance of animal welfare differs among developed and developing countries due to the level of economic
development, food security and education, as well as religious and cultural practices which presents challenges
for sound scientific research. Due to breeding goals in the past set on growth performance, traits such as fertil-
ity, welfare and health have been neglected. Fertility is the single most important trait in all livestock species.
Reduced fertility and lameness, claw health and mastitis results in unnecessary culling and reduced longevity.
Selection pressure for growth accompanied with inbreeding has resulted in a number of genetic defects in beef,
sheep and pigs. This review demonstrated the importance of inclusion of animal welfare concepts into breeding
objectives and selection strategies. Accurate phenotyping of welfare traits is a limiting factor in the implementa-
tion of mitigating strategies, which include diagnostic testing, control of inbreeding and genomic selection.
1 Introduction often have trade-offs and negative outcomes within the sys-
tem.
Quantitative selection practises have dominated the for-
The selection of farm animals commenced with the do- mulation of breeding objectives and selection over the past
mestication of the various livestock species approximately century. The development of animal recording systems re-
12 000 years ago, with the primary aim to benefit humankind sulted in a data-driven approach to formulation of breeding
in providing food, hides and fibres (Driscoll et al., 2009). A objectives. Garrick and Golden (2008) emphasised the po-
large variety of modern domesticated dairy and beef cattle, tential dangers of using only traits that are easily recorded
goat, sheep and pig breeds originated from their respective in selection programmes. These traits, such as growth traits,
ancestors, which included the auroch (Bos primigenius), be- have moderate to high heritability estimates and thus result in
zoar (Capra hircus), mouflon (Ovis aries) and wild boar (Sus relatively fast genetic progress. This approach has however
scrofa) (Diamond, 2002). neglected the traits not easily recorded or costly to record,
Over many centuries, livestock, pigs and poultry were including traits associated with health and animal welfare.
farmed under a wide range of environmental conditions in Some of the deleterious consequences in livestock produc-
different parts of the world. The arrival of various technolo- tion came about via direct selection, while others were unin-
gies resulted in modern production systems with advanced tended. The adverse effects on health and welfare traits have
housing, feeding and selection and breeding methodologies generally been more pronounced in intensive production sys-
for farm animal species that resulted in higher production tems, such as lameness in dairy cattle and pigs.
output and increased efficiency. However, biological and sci-
entific advances, such as selection for increased production,
Published by Copernicus Publications on behalf of the Leibniz Institute for Farm Animal Biology (FBN).178 E. van Marle-Köster and C. Visser: Consequences of selection for increased production
It is important to take note of the change in the interac- be defined by the animal’s individual perception of its own
tion between humans and their livestock over centuries. In physical and emotional state (Webster, 2013). The major-
ancient times and through the Middle Ages, humans lived ity of these all refer to physical health, environment and the
in a close relationship with their animals. Many communi- emotional and psychological state of the animal (Koknaroglu
ties throughout the world were dependant on some form of and Akunal, 2013). It could be argued that in the case of
agriculture, and the majority of agricultural production was farm animals, their physical health and production environ-
based on family farming. In developed countries, over many ment tend to be the primary focus with regard to animal
centuries, various factors contributed to a change in livestock welfare, as these aspects will have an influence on the pro-
farming, away from traditional family farms to larger com- duction outcomes and the profitability of the system. Animal
mercial production units with changed labour and manage- welfare should therefore be considered in selection to avoid
ment structures (Lowder et al., 2016). This has arguably also genetic defects or outcomes, which will result in unwanted
resulted in less close contact between farmers and their an- culling, pain or physiological stress. The physiological and
imals. In developing countries (as in Africa), a substantial behavioural aspects are more difficult to measure objectively
number of livestock are kept in communal and smallholder and often not considered in selection programmes.
systems where livestock serve several more purposes than In 1992, the Farm Animal Welfare Council compiled the
only being a food source (Mapiye et al., 2019). In these sys- “Five Freedoms of Animals” in an attempt to improve and
tems, different challenges are faced with regard to animal regulate the wellbeing of all farm animals (Manteca et al.,
health and welfare, such as the common practise of tether- 2012), which includes freedom from hunger and thirst, free-
ing (Mataveia et al., 2018). dom from discomfort, freedom from pain, injury or disease,
Selection of livestock in both intensive and extensive pro- freedom to express normal behaviour and freedom from fear
duction systems is under pressure to meet the increased de- and distress. These freedoms largely pertain to the living con-
mand for animal protein by a growing world population ditions and care of an animal. Furthermore, in 2008, the In-
which is estimated to reach 9 billion humans by 2050 (God- ternational Committee of the World Organisation for Animal
fray et al., 2010; Telugu et al., 2017). Furthermore, selection Health (also known by its French acronym: Office Interna-
programmes should address the reduction of the carbon foot- tional des Épizooties, OIE) passed a resolution appealing for
print and methane emissions and adhere to the sustainable affirmation of their standards of animal welfare as the global
development goals (SDGs). The consumer has also become reference standard for OIE members. Consequently, in 2012,
more conscious with regard to the origin of food production, the 178 members of the OIE adopted 10 “General Principles
standards and safety regulations. All these aspects are im- for the Welfare of Animals in Livestock Production Systems”
portant in the formulation of sustainable breeding objectives, (OIE, 2012), which are summarised in Table 1.
and specific attention should be given to traits related to ani- The relative importance of animal welfare differs among
mal welfare. The consumer has entered the debate on animal countries due to a variety of reasons that include the level of
welfare, often accompanied by emotion and little scientific economic development and food security, the level of edu-
reasoning (Buller et al., 2018). It is therefore the responsi- cation as well as religious and cultural practises (Wilkens et
bility of the animal science community to recognise welfare al., 2005; Sinclair et al., 2019). Perceptions of animal welfare
traits as important to the sustainability of the respective in- within the scientific community vary, which also presents
dustries. challenges for sound scientific research and maintaining ani-
In this review, we firstly provide context on animal wel- mal welfare as a scientific field (Korte et al., 2007; Von Key-
fare with regard to the current world perspective, followed serlingk and Hötzel, 2015).
by a discussion of the unintended consequences of selec- Policies and statutory regulations on animal welfare vary
tion on welfare and health traits with reference to extensive among countries. In many developed countries, animals are
and intensive production systems. Mitigating strategies are regarded as sentient creatures with a trend towards increased
reviewed with specific reference to welfare traits. anthropocentric thinking regarding the treatment of animals
(Korte et al., 2007). This may lead to an increased influence
of social and cultural opinions on legislation affecting animal
2 Defining welfare in livestock production and in effect may have a negative impact on the
global improvement of animal welfare.
In a broad sense, the word “welfare” is defined as the state in In Table 2, a list of non-governmental animal welfare or-
which an animal or human is content, safe and healthy (Ox- ganisations around the world is provided. This is not an ex-
ford University, 2009). The first definitions of animal wel- haustive list, but it demonstrates the recognition of animal
fare referred to the complete mental and physical health of welfare around the world.
the animal in harmony with its environment as well as free These organisations aim to protect a large number of
from suffering (Hughes, 1976; Carpenter, 1980). More recent animal species, including farm animals. Besides the non-
definitions state that the welfare of any sentient farmed ani- governmental organisations, there are international animal
mal, or animal used in the service of human beings, should welfare science organisations with the purpose to cover all
Arch. Anim. Breed., 64, 177–185, 2021 https://doi.org/10.5194/aab-64-177-2021E. van Marle-Köster and C. Visser: Consequences of selection for increased production 179
Table 1. Principles for the welfare of animals in livestock production (adapted from OIE, 2012).
Broad category Principle description
Genetic selection The effect on animal health, behaviour and temperament
External environment The influence on injuries and the transmission of diseases and parasites
Management The effect on resting, movement and the performance of natural behaviour of groups to min-
imise conflict and allow positive social contact
Housing The effects of air quality, temperature and humidity on animal health and comfort
Nutrition Ensuring access to feed and water suited to the animals/species needs
Veterinary and health Prevention and control of diseases and parasites, with humane euthanasia if treatment is not
care feasible or recovery is unlikely; prevention and management of pain
Emotional Creation of positive human–animal relationships
Human care and Ensuring adequate skills and knowledge among animal handlers
handling
Table 2. Examples of non-governmental animal welfare organisations around the world and the years in which they were founded.
Name of the organisation Year Reference (last access: 17 September 2020)
founded
World Animals Protection (WPSA) 1981 https://www.worldanimalprotection.or.ke/change-wspa
International fund for Animal Welfare (IFAW) 1969 https://www.ifaw.org/
Compassion in World Farming (CIWF) 1967 https://www.ciwf.com
Royal Society for the Prevention of Cruelty to Animals 1824 https://www.rspca.org.uk/
(RSPCA)
Eurogroup for Animal Welfare 1980 https://www.eurogroupforanimals.org/
National Council of Societies for the Prevention of Cru- 1955 https://nspca.co.za/
elty to Animals (NSPCA) South Africa
Japan Farm Animal Welfare Initiative 1973 https://www.esdaw.eu/society-and-animal-welfare-Japan
RSPC Australia 1981 https://www.rspca.org.au/
NSPCA New Zealand 1933 https://www.spca.nz/
aspects of ethology and interactions between humans and imal, which can be measured and included in selection pro-
animals, including farm animals (Bayvel and Cross, 2010). grammes.
It is important that animal geneticists and scientists consider The setting of breeding objectives, and thus selection
animal welfare traits in research efforts to ensure long-term progress, was historically driven by the economic value of
sustainable production outcomes. traits. Livestock industries aimed to make the most progress
in the production traits that would correspond to the high-
est profit (Miglior et al., 2017). This however was associ-
ated with a decline in other traits, mainly associated with re-
3 Unintended outcomes of selection
production and welfare. The focus of breeding objectives on
production in combination with a data-driven selection ap-
Welfare as mentioned in the section above is complex and
proach have resulted in unintended outcomes in a number
constitutes more than the emotional and physical wellbeing
of traits. In the seed stock industry, especially for dairy and
of the animal (Koknaroglu and Akunal, 2013). While emo-
pigs, genetic defects (e.g. CVM: complex vertebral malfor-
tional and physiological welfare traits remain challenging to
mation syndrome, BLAD: bovine leukocyte adhesion defi-
assess, there are a number of traits, which directly or indi-
ciency, MHS: malignant hypothermia-susceptible) have be-
rectly may influence the wellbeing and production of the an-
https://doi.org/10.5194/aab-64-177-2021 Arch. Anim. Breed., 64, 177–185, 2021180 E. van Marle-Köster and C. Visser: Consequences of selection for increased production come more prevalent where unintended inbreeding has been This is a complex trait in terms of recording and it is influ- practised with the extensive use of superior sires via artifi- enced by a range of physiological and management factors. cial insemination (AI). Although a number of countries have Selection for resistance to mastitis should result in resistance made a shift towards a more balanced approach regarding the to other diseases, as well as improved reproduction efficiency choice of selection objectives, it will take time to correct the and longer herd life of dairy cows (Martin et al., 2018). adverse effects of the previous strategy. In pig production systems, reproductive failures contribute The relationship between welfare and reproduction traits is to over 50 % of involuntary culling after the first farrow- complex and influenced by many factors. Although the rela- ing (Rauw, 1999). The use of AI in pig production sys- tionship between welfare and fertility is not linear (i.e. some tems has resulted in increased inbreeding rates, potentially animals can maintain high levels of productivity in the short causing inbreeding depression as well as increasing the fre- term while being compromised in terms of welfare aspects), quency of deleterious alleles (Andersson, 2013). These mu- a decrease in reproductive efficiency is associated with in- tations are usually present at low frequencies, with the al- creased stress (Ritter et al., 2019). The decline in fertility leles widespread throughout populations (Leroy, 2014). In- results in an increase in involuntary culling of the animals; tense selection and the use of AI have resulted in decreased therefore, it is a reasonable inference that fertility can be ac- effective population sizes that led to increased frequencies cepted as a welfare trait. of these deleterious alleles, resulting in increased prevalence Fertility is considered the single most important eco- of embryonic death caused by lethal mutations. This also af- nomic trait in both intensive and extensive production sys- fects fertility, lowering the reproductive performance in pig tems (Cammack et al., 2009; Walsh et al., 2011). Heritabil- populations (Derks et al., 2019). ity estimates for fertility traits are generally low due to the Lameness is a recognised problem in pigs (Le et al., 2015; high environmental variance. Intensive selection for produc- Nalon and Stevenson, 2019) and affects pig production on tion traits in the past has unintentionally led to selection for both a welfare and economic level. Le et al. (2015) reported impaired fertility, due to the unfavourable genetic correla- that boars with leg deformities and lameness showed a de- tions between these components. This decline in fertility is creased daily weight gain, resulting in decreased profitability the most pronounced in dairy cattle where milk yield, fat and efficiency. Sows with superior moving ability were likely yield and protein yield have been the major focus (Berry to show superior fertility, with larger litter sizes and shorter et al., 2014). In species such as dairy cattle, a vast amount between-parity intervals. Nalon and Stevenson (2019) added of research is available on the adverse effects of directional that locomotory disorders were the major cause of involun- selection for milk yield on cow fertility, health and welfare tary culling of healthy breeding sows. (Berry et al., 2003; Dillon et al., 2006; Roche, 2006; Zink Lameness has been also associated with claw quality in et al., 2011; Berry et al., 2014; Berry and Evans, 2014). beef cattle. Heritability estimates for claw traits remain low In the dairy industry, the use of AI has contributed to in- due to large environmental variances, and improved objec- breeding, resulting in a decline in overall fertility (Weigel, tive measurements are required for genetic evaluations. In a 2001). Genetic defects such as bovine leukocyte adhesion de- study on Angus cattle in the US, heritability estimates varied ficiency (BLAD), deficiency of uridine monophosphate syn- from 0.16 to 0.37 for different claw traits (Wang et al., 2017). thase (DUMPS) and complex vertebral malformation (CVM) In a South African study on Bonsmara cattle, the morpholog- are related to the overuse of specific superior sires and have ical and physiological characteristics of claws were studied, been well documented (Cole, 2015). indicating that tensile strength was influenced by bioregion. Dairy cow fertility is linked to cow lameness, which is It was also noted that the differences observed between front also related to claw health. Lameness is associated with pain and hind claws could be related to selection pressure on con- and stress, and the stress response is thought to have a neg- formation and accelerated growth (Van Marle-Köster et al., ative influence on the reproductive cycle, causing a decrease 2019). in number of eggs ovulated, as well as a decrease in fertili- Beef and sheep are raised mostly in extensive farming sys- sation rate (Fitzgerald et al., 2012). Lameness is associated tems with breeding objectives biased towards growth traits. with claw lesions (including non-infectious and infections Webb and Casey (2010) have reviewed the consequences disorders) which are mostly observed in cows housed in in- of prolonged selection for growth and growth efficiency on tensive production systems requiring higher management in- meat quality in detail. Selection emphasis on increased mus- puts. Claw defects such as cork screw claw, dermatitis, heel cularity and meat yield in beef cattle has inadvertently led horn erosion, sole ulcers and white line disorder have been to conditions such as double muscling with adverse effects reported to have a genetic basis (Heringstad et al., 2018) and on fertility (Fiems, 2012; Fiems and Ampe, 2014). The in- can be addressed in selection if accurately recorded on farm. activation of the myostatin (MSTN) gene causes an increase Solutions for improved claw welfare will require a multifac- in skeletal muscle weight, but at the same time, fertility de- torial approach. creases with an increase in dystocia and a reduction in calf Although health traits are beyond the scope of this review, survival (Arthur, 2004; Greger, 2011). A number of myo- mastitis must be mentioned as an important welfare trait. statin variants have been identified in beef cattle breeds using Arch. Anim. Breed., 64, 177–185, 2021 https://doi.org/10.5194/aab-64-177-2021
E. van Marle-Köster and C. Visser: Consequences of selection for increased production 181
genomics, and these are a reason for concern with regard to polymorphism (SNP) markers have provided numerous op-
animal welfare. portunities for studying genetic defects, developing commer-
Selection for accelerated growth also posed welfare prob- cial diagnostic tests and SNP-based parentage verification
lems in sheep and goat breeding. The callipyge gene in sheep panels, as well as routine genotyping for genomic selection
has been reported to cause a similar double-muscled pheno- and management of genomic inbreeding.
type in sheep to that seen in beef cattle; however, a study
done by Jackson et al. (1997) indicated that none of the ad- 4.1 Diagnostic testing
verse effects associated with double muscling in beef cat-
tle are seen in callipyge sheep. A decrease in terms of meat A large number of inherited disorders and familial traits in
tenderness have however been reported (Webb and Casey, animals (Lenffer et al., 2006) have been catalogued in the
2010). Online Mendelian Inheritance in Animals (OMIA). This cat-
In pigs, increased occurrences of metabolic diseases such alogue is continuously updated as new phenotypes are ob-
as porcine stress syndrome (PSS) and mulberry heart disease served and confirmed as inherited disorders. The origins of
have been linked to the genetic selection for increasingly fast most genetic defects remain uncertain. However, it is fair to
growing, lean animals, with large muscle blocks (Brambilla assume that some were fixated within certain breeds due to
et al., 2002). PSS is a major contributor to the occurrence linkage with economically important traits (thus as a conse-
of PSE (pale, soft and exudative meat) and has been linked quence of selection for production). Some were disseminated
to a recessive mutation on chromosome 6, characterised as throughout populations and breeds by the use of reproductive
a single-nucleotide substitution (T/C) in the gene coding for technologies such as AI and the overuse of some superior,
porcine calcium release, also referred to as the skeletal ryan- high-impact sires. SNP genotyping is an efficient methodol-
odine receptor 1 gene (RYR1 locus) (Ile et al., 2018). A num- ogy to identify carriers of genetic disorders (Ciepłoch et al.,
ber of other muscle defects (such as dark firm dry (DFD) syn- 2017). The causative mutations associated with a large num-
drome) have been reported in pigs (Webb and Casey, 2010). ber of defects are known and has been included on commer-
Few welfare studies on sheep and goats are available, cially available SNP arrays, to be identified during routine
which is largely attributed to the predominantly extensive genotyping.
systems in which they are produced (Sevi et al., 2009). Both Success stories of the eradication of genetic disorders us-
the ability of small stock to adapt to severe environments and ing DNA-based diagnostic testing include neuropathic hy-
their robustness are often exaggerated, leading to perceptions drocephalus found in Angus cattle populations (Teseling and
that inadequate management practises and unfavourable en- Parnell, 2013) and cardiomyopathy and woolly hair coat syn-
vironments do not have large influences on their health and drome (CWH) in Hereford cattle (Simpson et al., 2009). Sev-
welfare (Sevi et al., 2009). Husbandry practices that do im- eral other genetic disorders affecting beef cattle, sheep and
pact adversely on sheep include surgical mulesing and tail pigs can be identified using diagnostic tests. Some of these
docking. Mulesing is the removal of skin around the breech are available as either single gene-based tests or were added
area and is generally regarded as a painful procedure, as is to commercial SNP arrays. Table 3 presents a summary of
tail docking (Small et al., 2018). Mulesing has been banned genetic defects available on the SNP arrays for cattle, sheep
in some countries (i.e. New Zealand) but is still legal in and pigs, which can be requested in routine genotyping.
many others, including developed countries such as Aus- Genetic defects can be considered as part of animal wel-
tralia. There is a lack of information on welfare traits in fare as the birth of abnormal offspring should be prevented.
goats, especially in extensive production systems. Tether- Normal calving is regarded as a risk to welfare, due to the
ing is a common management practice in goat production in accompanying pain, stress and risk of dystocia (Ritter et al.,
Africa (Mataveia et al., 2018), and while it serves to protect 2019). Giving birth to an abnormal or stillborn calf is an un-
the animals from stock theft it also restricts their feeding and necessary additional trauma for the female animal as well as
often leads to both malnutrition and dehydration. a production loss for the farmer. Diagnostic testing can be
used to eliminate genetic defects from populations by identi-
fying phenotypically normal carrier animals (heterozygotes)
4 Mitigating strategies and preventing their mating with similar carriers. Genetic
disorders with significant economic impacts can therefore be
Genomic technology provides the opportunity to develop removed from the breeding population and culled to eradi-
mitigating strategies to manage the unintended negative con- cate the causative mutation completely.
sequences of directional selection. Several DNA markers
such as microsatellite markers have been used since the early 4.2 Management of inbreeding levels
1990s for DNA-based parentage verification and identifica-
tion of genetic defects. The completion of the full sequence The adverse effects of inbreeding have been well docu-
of the reference bovine genome in 2009 (Matukumalli et mented across all livestock species (Weigel, 2001; Anders-
al., 2009) and the subsequent discovery of single-nucleotide son, 2013). Incorrect and incomplete pedigree records how-
https://doi.org/10.5194/aab-64-177-2021 Arch. Anim. Breed., 64, 177–185, 2021182 E. van Marle-Köster and C. Visser: Consequences of selection for increased production
Table 3. Summary of the number of genetic defects included on SNP arrays for various livestock species.
Species SNP array No. Reference
defects
Beef Beef GGP HD150K array 48 Illumina (2020, 2015a)
Dairy Dairy GGP 50K array and dairy GGP HD150K array 18 Illumina (2020); Neogen Corporation (2020a)
Pigs Porcine 60KSNP array 5 Illumina (2015b); Neogen Corporation (2020b)
Sheep Ovine SNP50 beadchip 21 Neogen Corporation (2020c); https://SheepHapMap.org
(last access: 3 September 2020); Synnov (2016)
ever remain a main contributor to increased and unacceptable Genomic selection has also been applied in beef cattle
levels of inbreeding. One of the first and still one of the most worldwide where large numbers of beef cattle have been
important contributions of molecular genetics to livestock genotyped. Reliability of predictions improved with the use
breeding was the development of parentage verification pan- of genomic information and shows value, especially for the
els. These panels commonly consisted of a small number of difficult to measure traits such as feed efficiency (Hayes et
microsatellite markers but are now included as routine geno- al., 2013). Limited GS has been used for the improvement of
typing panels in SNP arrays. Correct parentage is further cru- health and welfare traits in beef cattle. Despite the potential
cial for accurately estimated genetic parameters (Visscher et of genomic information for selection of welfare and health
al., 2002; Pollak, 2005), breeding values and rate of genetic traits, accurate phenotypes remain a prerequisite for use of
progress (Van Eenennaam and Drake, 2012; Garritsen et al., the methodology (Coffey, 2020) and are currently the main
2015). In both the dairy and beef industries, breeders have challenge for implementation.
the opportunity to use SNP-based parentage testing and con- The field of genomics has shown significant progress in the
firm carrier animals with genetic defects as part of routine past decade, and genomic selection has become the method
genotyping. These technologies are still primarily available of choice to attain fast genetic progress. The benefits of this
to commercial farmers in developed countries and remain un- methodology are however limited to traits for which a large
affordable for the majority of developing countries. number of accurate phenotypes are available (Visser et al.,
2020). Phenotyping is thus the limiting factor for improving
4.3 Genomic selection reproductive and other welfare traits, such as clinical masti-
tis, claw lesions and hoof scoring. Countries with substantial
Genomic selection (GS) is the selection of animals for breed- phenotypes will reap the benefits of genomics, while in de-
ing based on both their estimated breeding values and a di- veloping countries such as South Africa, welfare traits are
rect genomic value, calculated from the combined effects not routinely recorded. Genomics will not improve animal
of markers spread across the genome (Goddard and Hayes, welfare if proper farm animal management practices, ani-
2009; Meuwissen et al., 2016). GS is most effective when mal recording and phenotyping are not in place. This must
applied to traits that are difficult to measure, expressed in be supported by an increased consciousness for animal wel-
only one gender or expressed late in life. This made GS an fare by animal scientists, farmers, industry and government.
attractive technology for the dairy cattle industry, which was
the first of the livestock industries to adopt the methodology.
5 Conclusions
The implementation of GS required training populations
and phenotypes. Due to extensive use of AI in the dairy in- The selection emphasis on production traits has clearly re-
dustry, they took the lead in implementation of GS. As in- sulted in a decrease in fitness traits including fertility, health
dicated by Berry et al. (2014), larger training populations and welfare of livestock. Selection for welfare and health
are required for lowly heritable traits. Studies have however traits is complex and most of the time not part of routine
proven the impact on the rate of genetic improvement in traits recording. It is necessary to find solutions to mitigate the un-
such as fertility, when genomic information was added in ge- intended consequences of directional selection and to attain
netic evaluations (Garcia-Ruiz et al., 2016; Ma et al., 2018). balanced breeding objectives that can be applied in sustain-
McNeel et al. (2017) demonstrated the value of genomic in- able breeding and efficient animal production. Genomics is
formation for prediction for a number of health and welfare one of the tools that can be used to select against genetic
traits, which included lameness and mastitis. Genomic in- defects, control inbreeding and add genomic information for
formation was added to the genetic evaluations to provide increased accuracy. This, however, should be reinforced by
a standard transmitting ability (STA) for the animals. The good management practices and an increased awareness of
authors however stressed the importance of available pheno- the importance of animal wellbeing.
types for these traits related to health and welfare.
Arch. Anim. Breed., 64, 177–185, 2021 https://doi.org/10.5194/aab-64-177-2021E. van Marle-Köster and C. Visser: Consequences of selection for increased production 183
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